Kurt A. Buhlmann - A biological inventory of eight in northwestern with conservation implications. Journal of and Studies 63(3): 91-98.

A BIOLOGICAL INVENTORY OF EIGHT CAVES IN NORTHWESTERN GEORGIA WITH CONSERVATION IMPLICATIONS

KURT A. BUHLMANN1 University of Georgia, Savannah River Ecology Laboratory, Aiken, SC 29802 USA

A 1995 biological inventory of 8 northwestern Georgia caves documented or re-confirmed the presence of 46 species of invertebrates, 35 considered troglobites or troglophiles. The study yielded new cave records for amphipods, isopods, diplurans, and carabid beetles. New state records for Georgia included a pselaphid beetle. Ten salamander species were in the 8 caves, including a true troglobite, the cave salamander. Two frog, 4 bat, and 1 rodent species were also documented. One cave contained a large colony of gray bats. For carabid beetles, leiodid beetles, and millipeds, the species differed between the caves of Pigeon and Lookout Mountain. Diplurans were absent from Lookout Mountain caves, yet were present in all Pigeon Mountain caves. A comparison between 1967 and 1995 inventories of Pettijohns Cave noted the absence of 2 species of drip pool amphipods from the latter. One cave had been contaminated by a petroleum spill and the expected aquatic fauna was not found. Further inventory work is suggested and the results should be applied to management strategies that provide for both biodiver- sity protection and recreational cave use.

Georgia is a cave-rich state, with most caves occurring in 29 July; Nash Waterfall Cave [NW] on 5 August; and Pigeon two distinct physiographic regions, the Cumberland Plateau Cave [PC] on 16 July (a) and 30 July (b). One cave was exam- and the southwestern Coastal Plain. Caves in the Cumberland ined in Chattanooga Valley in Walker County: Fricks Cave Plateau lie primarily in the counties of Dade, Walker, and [FC] on 16 September. Two caves were examined in Cloudland Chattooga. Canyon State Park at the base of Lookout Mountain in Dade A comprehensive inventory of Georgia caves was conduct- County: Case Cave [CC] and Sittons Cave [SI], both on 26 ed in 1967 (Holsinger & Peck 1971). Reeves et al. (2000) August (a) and 17 September (b) (Fig. 1). reported additional faunal records. Other scattered references The study followed methods used by Holsinger & Peck to the cave fauna of Georgia can be found in Loomis (1939: (1971) and Buhlmann (1992). Cave habitats sampled included millipeds), Hubricht (1943: amphipods), Chamberlin & streams, drip pools, phreatic lakes, decaying wood, cave walls, Hoffman (1958: millipeds), Hyman (1954: planarians), Barr and mud banks. Bait, usually liver cat food, was left in terres- (1965, 1981: beetles), Cooper (1968: salamanders), Holsinger trial habitats and checked on a return visit. Aquatic fauna in (1969, 1978: amphipods), Cooper & Iles (1971: fish), and Peck cave streams and drip pools were collected with small aquari- (1973: beetles). Dearolf (1953), Nicholas (1960), and um dip nets and suction tubes; baited containers were set in Holsinger & Culver (1988) provide checklists of species and deep pools. The first inventory of each cave included a search reviews of the regional biogeography of cave faunas. for all representative habitat types and collection of organisms. The goals of this study were to conduct biological invento- A second visit was made to some caves 1-2 weeks later to col- ries for cave-adapted species of northwest Georgia caves that lect specimens from the bait stations and revisit selected habi- were under state management responsibility and receive vary- tats. Four caves (PC, AS, CC, and SI) were each visited twice ing levels of recreational use, and/or were believed to have sig- during this study. Three caves (FC, EC, and NW) were visited nificant biodiversity and conservation value. Results may be once and 1 cave (PJ) was visited 3 times. useful in the development of cave management and conserva- Invertebrates were preserved in 70% ethanol for later iden- tion plans and will contribute to knowledge of the distribution tification. Only a minimum number of specimens were col- and biogeography of Georgia’s cave fauna. lected for species identification. Invertebrate taxa were sorted and shipped to recognized experts of the various taxonomic METHODS groups (see ACKNOWLEDGMENTS) where they remain or have been deposited in museum collections. Data were collected on Between 15 July and 14 October 1995, biological invento- habitats, species abundance, and human disturbance. Simply ries were conducted in 8 caves within northwestern Georgia’s documenting presence or absence was the greatest level of Cumberland Plateau. Five caves were examined on Pigeon detail attainable for many species. Nomenclature used in the Mountain in Walker County: Pettijohns Cave [PJ] on 15 July (a), 5 August (b), and 14 October (c); Anderson Springs Cave 1 Current address: Conservation International, Center for Applied Biodiversity Science, [AS] on 30 July (a) and 6 August (b); Ellisons Cave [EC] on 1919 M Street, NW, Suite 600, Washington DC 20036 USA [email protected]

Copyright © 2001 by The National Speleological Society Journal of Cave and Karst Studies, December 2001 • 91 BIOLOGICAL INVENTORY OF EIGHT CAVES IN NORTHWESTERN GEORGIA

RESULTS

Class Turbellaria (flatworms), Order Tricladida, Family Kenkiidae: Sphalloplana sp. (TB): PJ(a). One specimen was collected in a pool at the base of a waterfall. The specimen may be Sphalloplana georgiana (Hyman) which was described from a cave in Dade Co., GA (Hyman 1954). Class , Order Isopoda (isopods), Family Asellidae: Caecidotea cyrtorhynchus Fleming and Steeves (TB): PJ(a); NW; AS(a,b). Individuals were especially abundant in the out-flowing, permanent stream in AS. Caecidotea sp. (undescribed or possibly Caecidotea catachaetus Fleming and Steeves (TB): EC; SI(b). Specimens were collected in a pool in the drying cave stream in SI. Two species of aquatic troglobitic isopods are currently known from GA, C. cyrtorhynchus and C. catachaetus (Fleming & Steeves 1972). Lirceus sp. (TP): NW. Specimens were collected from drip pools. Family Ligiidae: Ligidium elrodi Packard (TP): EC; PC(a). This terrestrial species was found around the entrance to NW. L. elrodi are perhaps becoming adapted to cave life (Schultz 1970). They inhabit wet, damp places, including the margins of springs and streams. The specimens found at NW are L. elrodi hancockensis (Schultz) (J. Lewis, pers. comm.). Family Trichoniscidae: Amerigoniscus sp. (possibly undescribed, TB): CC(a); SI(a). Prepared dissec- tions of this terrestrial isopod did not match known species from the area (J. Lewis, pers. comm.). Miktoniscus sp. (probably Miktoniscus alabamensis Muchmore, TP): PC(a). M. alabamensis is a widely ranging species known from AL, FL, and VA (Muchmore 1964). Order (amphipods), Family : Crangonyx antennatus Packard (TB): FC; SI(b); PJ(a); AS(a,b). Specimens in FC and AS were collected from streams with cobble bottom. Specimens from SI and PJ were collected from mud-bottomed streams. C. antennatus is wide- spread, and recorded from caves in southwest VA, eastern TN, northwestern GA, and northern (Holsinger & Culver 1988). minutus Holsinger (TB): NW. One specimen was collected in a drip pool. The species was previously collected from mud-bottomed drip pools Figure 1. General location of eight caves inventoried dur- in PJ (Holsinger 1978). ing the study (in ellipses): A. - Cloudland Canyon State Order (), Family : Park (Case Cave [CC] and Sittons Cave [SI]); B. - sp. (TX): EC; SI(a,b). Several specimens were observed in cave streams, but no troglomorphic species were found. Chattanooga Valley (Fricks Cave [FC]); C. - Pigeon Class Arachnida, Order Pseudoscorpiones (pseudoscorpions) Mountain (Pettijohns Cave [PJ], Anderson Springs Cave No specimens were found. Troglobitic species of pseudoscorpions have been [AS], Ellisons Cave [EC], Nash Waterfall Cave [NW], and previously found in Chattooga, Dade, and Walker County, GA caves Pigeon Cave [PC]). (Holsinger & Peck 1971), but not in any of the caves investigated during this study. Order Acari (parasitic mites) Results follows Peck (1998), unless other literature was unknown sp. (AC): PC(a); CC(a); NW; FC. Several genera of mites were col- required. lected, but all are believed to represent surface species carried into caves by Cave organisms were classified into one of four categories bats. Holsinger & Peck (1971) reported a cave-adapted mite from PJ. Order Opiliones (harvestmen), Family Phalangodidae: of cavernicoles (Barr 1963a, 1968). Troglobites (TB) are oblig- Bishopella sp. (TP): PC(a); EC; NW; CC(a,b); SI(b); FC; PJ(c). Adult and atory cave species with morphological adaptations for the cave juvenile specimens of Bishopella were collected in seven caves during this environment and do not exist in surface environments. study and Reeves et al. (2000) subsequently collected Bishopella in AS. Troglophiles (TP) may frequent caves and are capable of com- Bishopella have -developed eyes and have been collected occasionally from epigean habitats (Goodnight & Goodnight 1960). The Bishopella pleting all stages of their life cycle within a cave, but also occur likely includes several species that have been reported as Phalangodes lacin- in surface habitats. Trogloxenes (TX) are species that may use iosa (Crosby & Bishop) from northwestern GA caves, including SI, PJ, and cave environments seasonally or for portions of their life cycle, also northern AL and central TN caves (Holsinger & Peck 1971). yet must also be in association with surface environments. Order Araneae (), Family Agelenidae: Calymmaria sp. (TP): NW. Adults of both sexes were collected and may rep- Accidentals (AC) are those species that are often found in resent Calymmaria cavicola (Banks), a species described from other Walker caves but generally exist there only temporarily. Results focus Co. caves, as well as caves in TN and AL (Heiss 1982). on troglobitic and troglophilic species, with some exceptions. Family Amaurobiidae: Coras juvenilis Keyserling (TX?): FC. Female and immature specimens were collected and apparently represent a state record for the species (M. Draney, pers. comm.). C. juvenilis has not been reported from caves, although con- geners (Coras cavernorum Barrows from NC and Coras taugynus Chamberlin from AL) have been found in caves (Muma 1946).

92 • Journal of Cave and Karst Studies, December 2001 BUHLMANN

Family Leptonetidae: Typhlobius ?caecus Bollman (TX?): FC. The posterior end of a female litho- Leptoneta sp. (TP): PJ(a,c). Immature specimens were collected. Leptoneta is biid centipede was found. An intact adult male would be very valuable since represented in the western and southeastern U.S. by ~35 species; many are records for cave-dwelling centipedes are scarce and their status as troglobites cave dwellers (Roth 1993). Holsinger & Peck (1971) collected an undescribed has not been determined (R.L. Hoffman, pers. comm.). Leptoneta in PJ. Class Insecta, Order Collembola (springtails), Family Entomobryidae: Family Linyphiidae: Pseudosinella christianseni Salmon (TB): PJ(a,b,c); PC(a); EC; NW; AS(a,b); Phanetta subterranea (Emerton) (TB): PC(a); FC; SI(a). Phanetta is a mono- FC; CC(a,b); SI(a,b). Springtails were found in every cave visited and were typic genus (Millidge, 1984). It has been reported from caves in PA, MD, WV, often abundant around damp and decaying wood. P. christianseni is an artifi- VA, TN, KY, IL, IN, and AL (Peck & Lewis 1978; Roth 1988). P. subterranea cial taxon that will eventually be split into a series of species, all of which are is known from GA caves in Floyd, Walker, and Dade Co., including SI highly troglomorphic (K. Christiansen, pers. comm.). (Holsinger & Peck 1971). Pseudosinella n. sp. (TP): EC; NW; PC(a); FC. Pseudosinella n. sp. is not Family Mysmenidae: troglomorphic, but all of the P. n. sp. specimens collected are the same species Maymena ambita (Barrows) (TX ): EC. This is not a strict cave-dweller. (K. Christiansen, pers. comm.). It has been found in caves in AL, TN, and KY (Gertsch 1960), as well as one Family Isotomidae: other Walker Co., GA cave (Holsinger & Peck 1971). Folsomia candida Willem (TP): PJ(b). F. candida is an opportunistic Family Nesticidae: troglophile and has been recorded from caves over a wide area of the United Nesticus sp. 1 (TB): PC(a); AS(a). This was a small, eyeless purple spider (M. States (Christiansen 1982). Draney, pers. comm.). More specimens are needed. Family Tomoceridae: Nesticus sp. 2 (TP): CC(a,b). Several species of Nesticus with restricted ranges Tomocerus bidentatus Folsom (TP): EC; NW; PC(a); CC(b). T. bidentatus is a are known from caves in TN and VA (Holsinger & Culver 1988; Coyle & common and widespread collembolan in caves in the eastern U.S. It is lightly McGarity 1991) and recent genetic analyses indicate considerable genetic pigmented with small eyes (Christiansen 1982; Holsinger & Culver 1988). divergences among recognized taxa within the genus Nesticus (Hedin 1997). Order Diplura (diplurans or bristletails), Family Campodeidae: Distributional data from Hedin (1997) and a report by Reeves et al. (2000) of Litocampa sp. “P” (TB): PJ(a,c); EC; AS(a,b); PC(a); NW; FC. Diplurans Nesticus georgia in Sittons Cave (SI) indicate that these specimens from CC were found on undisturbed mud banks. Litocampa sp. “P” was previously may represent N. georgia Gertsch. known only from PJ and a cave in Chattooga Co., GA. This study found five Family : new cave localities for this possible Georgia endemic (L. Ferguson, pers. Liocranoides sp. (TP): PJ(a,c); AS(a). The specimens are likely Liocranoides comm.). Litocampa are not known from CC and SI. Three Litocampa species gertschi Platnick (Platnick 1999). Specimens of this spider previously report- are known from GA caves (Ferguson 1981). The species designation of “P” ed from several Walker County caves (Holsinger & Peck 1971) as corresponds with a coding system for unnamed species (L. Ferguson, pers. Liocranoides unicolor Keyserling, are likely L. gertschi. comm.). Family Tetragnathidae: Order Orthoptera, Family Rhaphidophoridae (cave and camel crickets): Meta menardi (Latreille) (TP): NW; PC(a); FC. This orb weaver was fre- Ceuthophilus gracilipes (Haldeman) (TX): The common “camel” cricket was quently seen in some cave entrances. It is also found under bridges, in damp found in the entrances of all 8 caves. Hundreds of camel crickets were ravines, and hollow logs (Marusik & Koponen 1992; M. Draney, pers. comm.). observed under a ledge at the entrance to PC(b). C. gracilipes occurs through- Reeves et al. (2000) reported Meta from FC and SI. out the eastern in forested habitats as well as cave entrances and Family Theridiidae: no systematic effort was made to collect them. Achaearanea porteri (Banks) (TX): FC. This spider has been recorded from Euhadenocecus puteanus Scudder (TP): PC(a); EC; AS(a); FC; CC(a). E. MD to TX (M. Draney, pers. comm.). The previous GA records are not from puteanus is widely distributed throughout the Appalachian region and the caves (Levi 1955). Interior Low Plateaus and is closely associated with cave environments Class Diplopoda (millipeds) Order Spirostrepida Family Cambalidae: (Holsinger & Culver 1988). It has been recorded from GA caves in Dade, Cambala hubrichti Hoffman (TX): FC. These large pinkish-gray millipeds Walker, and Polk Co. (Holsinger & Peck 1971). were abundant on gray bat (Myotis grisescens) guano in FC. C. hubrichti is Order Diptera (flies), Family Heleomyzidae: also known from caves in western SC (Chamberlin & Hoffman 1958). undetermined Heleomyzidae (TP): These flies were observed on the ceilings Order Chordeumatida Family Cleidogonidae: and damp walls near the entrance zones of all caves. No systematic effort was Pseudotremia eburnea Loomis (TB): CC(a,b); PJ(b); EC. P. eburnea was made to collect cave flies. described from a cave in Dade County, GA (Loomis 1939; Chamberlin 1946), Order Coleoptera (beetles), Family Carabidae (ground beetles): and was reported from CC by Shear (1972) and from PJ by Holsinger & Peck Pseudanophthalmus georgiae Barr (TB): PJ(a,c); EC. One of the PJ specimens (1971). Reeves et al. (2000) reported P. eburnea from several additional collected was white. Holsinger & Peck (1971) collected this species, unde- Georgia caves. scribed at that time, in PJ and Mountain Cove Farm Cave. It is also known Pseudotremia sp. 1 (TB): NW. These female specimens probably represent P. from Blowing Springs Cave (Barr 1981). The EC specimen represents a new eburnea, but could be a different species (R.L. Hoffman, pers. comm.). cave record for P. georgiae. Pseudotremia sp. 2 (TB): SI(a). Holsinger & Peck (1971) also collected an Pseudanophthalmus fulleri Valentine (TB): SI(a). P. fulleri was previously undescribed Pseudotremia from SI. known from SI (Barr 1981). It was also collected in several Dade and Walker Pseudotremia aeacus Shear (TB): PC(a,b). P. aeacus was described from a Co. caves by Holsinger & Peck (1971) and Reeves et al. (2000) and is also cave in Dade Co., GA (Shear 1972); the PC specimens may represent a new known from caves in Hamilton Co., TN. subspecies (R.L. Hoffman, pers. comm.). Family Leiodidae (scavenger beetles): Family Trichopetalidae: Ptomaphagus fiskei Peck (TB): PJ(a,b,c); EC; PC(b); AS(a,b). P. fiskei was Scoterpes austrinus Loomis (TB): PJ(c); EC; NW; AS(a,b); SI(b). S. austrinus especially abundant at bait stations in PJ. P. fiskei appears to be restricted to was previously found in several caves, including PJ (Holsinger & Peck 1971), Pigeon Mountain. According to Peck (1973), no Ptomaphagus are known from and as well as caves in DeKalb Co., AL (Loomis 1943). caves in the valley between Pigeon and Lookout Mountains. No Ptomaphagus Scoterpes sp. (TB): NW; PC(a,b); FC; CC(b). These specimens are perhaps S. were found in FC. austrinus or represent a new species. Holsinger & Peck (1971) suggested that Ptomaphagus whiteselli Barr (TB): CC(a,b); SI(a,b). P. whiteselli was previ- at least two subspecies (or species) of Scoterpes exist in GA caves. Additional ously known from SI (Barr 1963b) and CC (Peck 1973). It is the only specimens, specifically adult males, are required (R.L. Hoffman, pers. comm.). Ptomaphagus known from caves between Sand and Lookout Mountains in Order Polydesmida Family Xystodesmidae: DeKalb Co., AL and Dade Co., GA (Peck 1973). Cherokia georgiana Bollman (AC): PC(a). Specimens were collected at the Family Pselaphidae (pselaphid beetles): entrance to PC. C. georgiana is not usually considered a cave inhabitant. Batrisodes sp. (TX): PC(a). The specimens may be Batrisodes globosus Class Chilopoda (centipedes) Order Lithobiomorpha Family Lithobiidae: (LeConte), a widely distributed species in eastern and record-

Journal of Cave and Karst Studies, December 2001 • 93 BIOLOGICAL INVENTORY OF EIGHT CAVES IN NORTHWESTERN GEORGIA

ed from caves in AL, GA, and VA (Barr 1964; Park 1947; Holsinger & Peck petraeus was discovered, representing the northernmost locality for this 1971). Pigeon Mountain endemic. The locality is not reported here in order to protect Speleochus sp. (TB): PC(a). PC produced the only specimen of this unde- the population. scribed species known from GA. The genus Speleochus is only known from Desmognathus fuscus (Green) (northern dusky salamander, AC): PC(a); EC; central TN, northeast AL, and Walker Co., GA (T. Barr, pers. comm.). Species NW. At PC, several individuals were found near the entrance under rocks in in this genus seem rare and they appear to be most abundant during cold, wet the in-flowing cave stream. 2 salamanders were observed at the entrance to EC weather (Jan-Feb) when they may be more easily observed because their inter- and 1 was observed at the entrance to NW. stitial habitats are flooded. (T. Barr, pers. comm.). Pseudotriton ruber (Latreille) (red salamander, AC): PC(b); AS(a,b). In PC, 2 Family Staphylinidae (rove beetles) (TX): adult red salamanders were seen in the cave stream. In AS(a), 1 adult was Staphylinid beetles were collected in PJ, NW, PC, and FC and are in the pos- observed and in AS(b), 2 adults were observed. Red salamanders are not usu- session of T. Barr. Staphylinid beetles were commonly found at bait traps. ally considered cave associates. None of the species known are restricted to caves (Holsinger & Culver 1988). Class Mammalia, Order Chiroptera ( Bats), Family Vespertilionidae: Class Osteichthyes, Order Perciformes, Family Cottidae: Pipistrellus subflavus (Cuvier) (eastern pipistrelle, TX): PJ(a), 1 observed; Cottus bairdi (mottled sculpin, TX): FC. Several sculpins were observed in the PC(a), 1 obs; EC, 68 obs; AS(a), 36 obs; FC, several obs; SI(a), 1 obs. and FC stream; 1 was collected and dissected, but no food items were found in the SI(b), 5 obs. Eastern pipistrelles are the most common cave bat in eastern stomach. North America (Harvey 1992). They usually hang singly in the warmer parts Class Amphibia, Order Anura (frogs and toads), Family Ranidae: of the cave. Martin & Bearden (1990) reported P. subflavus to be abundant in Rana palustris LeConte (pickerel frog, TX): EC; AS(a); NW. 5 frogs were both EC and AS during inventories conducted Jan 1989-Feb 1990. observed inside the entrance to EC; 1 frog each was found in AS and NW. Myotis lucifugus (LeConte) (little brown bat, TX): EC, 1 was observed. Pickerel frogs have been commonly observed inside VA caves (K.A. Myotis grisescens (Howell) (gray bat, TX): FC; SI(b). At FC, a bachelor Buhlmann, pers. obs.). colony of 10,000-15,000 individuals was observed. At SI(b), 4 gray bats were Rana clamitans Latreille (green frog, AC): EC; NW. 1 frog each was found observed, 2 were females. Large piles of guano in FC indicate that the cave has near the entrances in EC and NW. Unlike pickerel frogs, green frogs are not been used by gray bats for many years. FC is relatively pristine, and has been commonly reported from caves. closed to for nearly 30 years (A. Padgett, pers. comm.). Gray bats are Order Caudata (salamanders) Family Plethodontidae: listed as federally Endangered and ~95% of the global population of gray bats Gyrinophilus porphyriticus (Green) (northern spring salamander, TP): PJ(a); hibernates in 8 caves in TN, MO, KY, AL, and AR (Harvey 1992). The 4 gray EC; AS(a,b); PC(a,b). The PJ specimen was a pale larva that was re-absorbing bats observed in SI likely represent individuals on migration (C. Hobson, pers. its gills. 1 adult individual was observed in the cave stream in EC. An adult and obs.). However, a gray bat stain on the ceiling indicates that SI may have his- a larva were each observed in AS. In PC(a) 4 larval individuals and 1 adult torically harbored a gray bat maternity colony. were observed in the cave stream. Also in PC(b), 2 very large larval individu- Myotis septentrionalis (Trouessart) (northern long-eared bat, TX): CC(a), 1 als were collected at the back of the cave, 370 m from the entrance, in the ter- long-eared bat was observed; SI(b), 3 male long-eared bats were observed. minal siphon. These very pale larvae with seemingly reduced eyes were Order Rodentia (gnawing mammals), Family Cricetidae: deposited in the U.S. National Museum of Natural History, Washington D.C. Neotoma floridana (Ord) (eastern woodrat, TX): Although no woodrats were (USNM 497685, 497686). observed during the study, their presence within each cave was recorded as Gyrinophilus palleucus McGrady (Tennessee cave salamander, TB): FC. 4 lar- active or historic after investigating droppings and nests: PJ, historic; EC, val individuals were observed in the cave’s out-flowing stream and 1 specimen active; PC, active; NW, active; CC, historic. No woodrat sign was found in AS, was collected (USNM 497687). The nearest AL records for G. palleucus are FC, and SI. Woodrat populations have been declining in the northeastern ~16 km west of Chattanooga Valley on the AL/GA line near Rising Fawn, GA United States and the species is monitored by several state Natural Heritage (Godwin 1995). Cooper (1968) reported a GA specimen but the locality is Programs (C. Hobson, pers. comm.). uncertain. The FC specimens confirm the presence of G. palleucus in GA (Buhlmann & Wynn 1996). COMPARISON OF THE 1967 AND 1995 Eurycea lucifuga Rafinesque (cave salamander, TP): All individuals reported were observed and not collected: PJ(a), 3 adults; PC(a), 2 adults and several STUDIES OF PETTIJOHNS CAV E larvae in the cave stream; EC, 1 adult and several larvae in cave stream pools; AS(a), 9 adults and AS(b), 5 adults; NW, 5 adults; CC(a,b), 1 adult each; SI(a), The most comprehensive Georgia cave study was conduct- 4 adults and SI(b), 1 adult. Cave salamanders are often encountered in caves ed in 1967 and one of the caves studied was PJ (Holsinger & throughout the Appalachian and Cumberland regions. Eurycea longicauda (Green) (longtail salamander, TX): All individuals report- Peck 1971). A comparison of the 1967 and 1995 studies found ed were observed, not collected: AS(a), 4 adults and AS (b), 5 adults; FC, 1 some faunal differences in PJ, primarily with regards to miss- adult. Longtail salamanders, although closely related to cave salamanders (E. ing aquatic fauna in 1995 (Table 1). Stygobromus minutus and lucifuga), are occasionally, although not commonly found in caves. S. dicksoni, amphipods that inhabit drip pools (Holsinger Plethodon glutinosus (Green) (slimy salamander, TX): EC; PJ(b); AS(a,b); NW; CC(a). In EC, 64 slimy salamanders were seen inside the Historic 1978) were not found in 1995, nor were they found by Reeves Entrance in the first 50 m, and an additional 81 salamanders were seen et al. (2000). between 51-60 m of this entrance. 20 more were observed inside the New The 1967 study also reported terrestrial isopods, pseu- Entrance to EC, totaling 165. 10 slimy salamanders were seen at the entrance doscorpions, and cave-adapted mites, none of which were to PJ(b); 9 were seen in AS(a) and 4 were observed in AS(b); 2 were observed in NW; and 1 was observed in CC(a). The large numbers of P. glutinosus in the found by the 1995 study. Pseudoscorpions and mites are often entrance to Ellisons Cave seemed unusual and warrant further study. difficult to detect in caves, yet terrestrial isopods Plethodon dorsalis (Cope) (zigzag salamander, AC): PJ(c). 1 specimen was (Amerigoniscus) should have been found by the methods used observed 10 m inside the entrance. in 1995. The milliped, Cambala hubrichti, collected in 1967, Plethodon serratus Grobman (southern redback salamander, AC): AS(b). 1 adult was observed at the entrance. was not found in 1995. Interestingly, this study found C. Plethodon petraeus (Wynn, Highton, and Jacobs) (Pigeon Mountain salaman- hubrichti abundantly in FC where it appeared to be associated der, TX): PJ(b). 1 very large adult was observed at the entrance. Plethodon with gray bat (M. grisescens) guano deposits. The 1995 study petraeus was expected in larger numbers around the entrance to PJ. It was documented an additional species each of troglophilic spider, apparently very abundant around the PJ entrance when it was first discovered (Wynn et al. 1988), but may since have been collected for the pet trade (A. collembolan, and beetle from PJ, although all are known from Wynn, pers. comm.). Incidental to the cave inventory, another location for P. other caves.

94 • Journal of Cave and Karst Studies, December 2001 BUHLMANN

TABLE 1. A comparison of invertebrate troglobites (TB) caves studied in Chattanooga Valley or on Lookout Mountain and troglophiles (TP) collected from Pettijohns Cave, (PJ=17, EC=17, NW=17, PC=17, AS=14, FC=13, CC=13, and Walker County, Georgia, during June 1967 (Holsinger & SI=11). The faunas of the Pigeon Mountain caves were simi- Peck 1971) and this study, July-October 1995. Names used lar, although several species were found in only 1 or 2 caves. by Holsinger & Peck are given in parentheses. However, faunal composition differed between the caves of Pigeon and Lookout Mountains. The terrestrial isopods SPECIES Holsinger & Peck, 1971 This Study (Amerigoniscus sp.) were found only in the 2 Lookout Mountain caves. Conversely, diplurans (Litocampa sp. “P”) Sphalloplana sp. X Caecidotea cyrtorhynchus X (as Asellus sp.) X were found in all Pigeon Mountain caves and FC, but not the Amerigoniscus sp. X (as Caucasonethes sp.) Lookout Mountain caves. The leiodid beetles were represented Crangonyx antennatus XX Stygobromus minutus X (as Stygobromus sp.) by 2 species with P. fiskei occurring in 4 of 5 Pigeon Mountain Stygobromus dicksoni X (as Stygobromus sp.) caves, while P. whiteselli was found in both Lookout Mountain Microcreagris sp. X caves. The troglobitic carabid beetles showed a similar 2 Rhagidia sp. X Bishopella sp. X (as Phalangodes laciniosa)X species distribution, with P. georgiae occurring in 2 Pigeon Leptoneta sp. X X Mountain caves, and P. fulleri being found in CC. The appar- Liocranoides sp. X Pseudotremia eburnea XXent absence of both Pseudanophthalmus and Ptomaphagus Scoterpes austrinus XXfrom FC is interesting in terms of biogeography. Peck (1973) Cambala hubrichti X (as Cambala minor) Pseudosinella christianseni X (as Pseudosinella hirsuta)Xhad previously hypothesized that the cave-adapted beetles may Pseudosinella n. sp . X be restricted to the flat-bedded limestones of the Appalachian Folsomia candida X Plateau. Peck’s hypothesis would explain the absence of bee- Litocampa sp. P X (as Plusiocampa sp.) X unidentified Heleomyzidae X (as Amoebalaria defressa)Xtles in FC and is also supported because different leiodid and Pseudanophthalmus georgiae X (as Pseudanophthalmus sp.) X carabid beetles are found on Lookout and Pigeon Mountains. Ptomaphagus fiskei X (as Ptomaphagus sp.) X The habitats within each cave may determine the faunal TOTALS 17 15 composition. AS has an out-flowing cave stream that originates within Pigeon Mountain and contained large populations of aquatic isopods (C. cyrtorhynchus), as well as some DISCUSSION amphipods (C. antennatus). Similarly, SI contained a large out-flowing stream and stygobitic were also noted, SPECIES AND DISTRIBUTION although in smaller numbers. Few aquatic cave organisms A total of 46 invertebrate taxa were identified during the were found in caves that contained in-flowing streams, notably study. Of those, 21 are considered troglobites, 14 are EC and PC. Small drip pools were noted in NW and contained troglophiles, and the remainder were classified as trogloxenes the amphipod, S. minutus, known previously only from PJ, as (9) or accidentals (2). Harvestmen (Bishopella sp.) and spring- well as an undescribed isopod (Lirceus sp.), and illustrate the tails (Pseudosinella sp.) are believed to represent several dif- importance of these habitats. CC has deep connections to ferent species and further taxonomic work is needed. Other phreatic water and a 2.5 m change in water level was noted collected specimens, mainly non-troglobitic, have been given between the two visits. The phreatic lakes of CC were previ- to others who are working with the material (W. Reeves, ously known to contain amphipods, isopods, and cave fish (A. Clemson Univ.). The pselaphid beetle (Speleochus sp.) was Padgett, pers. comm.). Limited observations indicate that cave found only in Pigeon Cave and represents a new, undescribed fish and Tennessee cave salamanders are most likely to be species for Georgia. The amphipod, Stygobromus minutus col- found in Georgia caves that have connections to phreatic lected in NW, was previously known only from PJ (Holsinger waters. Cave stream mud banks represent habitat for many of 1978), but was not found there during this study. Possible new the terrestrial invertebrates. Flooding streams deposit organic species of millipeds were also collected in NW and material that is scavenged by leiodid beetles, diplurans, mil- Pseudotremia sp. 2 is a possible endemic to SI. An undescribed lipeds and collembolans. The untrampled mud banks in AS and springtail (Pseudosinella) was collected in three of the Pigeon FC contained the largest numbers and diversity of terrestrial Mountain caves, as well as in FC. The dipluran (Litocampa sp. cave fauna. “P”) was previously known only from PJ, but was found in all Pigeon Mountain caves studied, as well as FC. EC yielded a MANAGEMENT IMPLICATIONS new cave record for the beetle, P. georgiae. For vertebrate All 8 caves inventoried are in public ownership or protect- species, 10 salamanders, 2 frogs, 1 fish, 4 bats, and 1 rodent ed by private owners. Four of the Pigeon Mountain caves are were encountered. In FC, the Tennessee cave salamander on the Crockford-Pigeon Mountain Wildlife Management Area (Gyrinophilus palleucus) was re-confirmed as a component of (CPMWMA) and 1 is immediately adjacent in private owner- Georgia’s fauna and a large population of federally endangered ship. Cloudland Canyon State Park (CCSP) on Lookout gray bats (Myotis grisescens) was found. Mountain contains CC and SI. The Georgia Department of Caves on Pigeon Mountain contained more species than the Natural Resources owns both CPMWMA and CCSP and The

Journal of Cave and Karst Studies, December 2001 • 95 BIOLOGICAL INVENTORY OF EIGHT CAVES IN NORTHWESTERN GEORGIA

Southeastern Cave Conservancy owns FC. Therefore, the Further inventory and research should address the protection of opportunity exists to effectively manage all of these caves for the landscape around caves (e.g., Aley & Aley 1991) in order both biodiversity protection and recreation. to protect cave habitats and fauna (e.g., Fong 1995; Jacobson PJ is probably the most frequently visited recreational cave 1995). Caves are unique natural habitats and contain endemic in Georgia. As many as 350 people have visited the cave in one and rare species. Further study of the biota and ecology of month and as many as 75 on a given weekend (Georgia DNR, Georgia caves is therefore warranted. unpubl. data). There are many passages and although all receive human traffic, some are more heavily used than others. ACKNOWLEDGMENTS The greatest numbers of collembolans and diplurans were found in PJ rooms and passages that contained signs of heavy For providing critical review of the manuscript I thank J.R. human use and garbage. It is unknown if the presence of addi- Holsinger, S.J. Taylor, J.J. Lewis, J.B. Jensen, and two anony- tional food resources results in increased populations of cave mous reviewers. For identifying specimens I thank: T.C. Barr, fauna or if it serves to attract and concentrate cave organisms. K.A. Christiansen, D.A. Crossley, M. Draney, L.R. Ferguson, If it serves to concentrate organisms, then are populations ulti- R.L. Hoffman, J.R. Holsinger, J.J. Lewis, S.B. Peck, D. Ubick, mately reduced by trampling? These questions could be and A.H. Wynn. Cavers and biologists who participated on one addressed with future research. Few bats were observed in PJ or more field trips included: C. Anderson, J. Bearden, M. and more Pigeon Mountain salamanders were expected based Branch, L. Clarke, J.P. Demuth, G. Eck, C.A. Hobson, N. on discussions with biologists who had observed them in the Holcomb, T. Holcomb, K. Huffines, R. Kappel, J.A. Ott, J. past. Overall, PJ contained a great diversity of cave life, par- Ozier, A. Padgett, K. Padgett, M.A. Pilgrim, J. Respess, D. ticularly terrestrial species. Aquatic organisms seemed rare in Sorrell, S. Sotona, D. Wohl, G. Yanochko, H. Young, and S. PJ, perhaps a result of trampling in drip pools and streams. No Young. A. DeBiase and B. Taylor of SREL allowed use of amphipods (Stygobromus sp.) were found during this study in their laboratory for sorting specimens. A. Padgett, N. PJ, yet two drip pool species, S. dicksoni and S. minutus were Holcomb, and J. Bearden of the Georgia Department of recorded previously (Holsinger & Peck 1971). However, not Natural Resources provided logistical support. Financial sup- all stream passages were explored and refugia may exist, par- port was provided by The Athens Speleological Society, The ticularly in the less traveled passages. Protection of certain Georgia Speleological Survey, The Dogwood City , The passages might provide refugia for cave organisms while con- Clayton County Cavers, The Chattanooga Grotto, The tinuing to provide recreational opportunities for cavers. Southeastern Region of the NSS, and the Georgia DNR. Stygobromus dicksoni is also known from several caves in Manuscript preparation was supported by the University of northeastern AL and northwestern GA (Holsinger 1978; Georgia’s Savannah River Ecology Lab through Contract #DE- Reeves et al. 2000). FC09-96SR18546 with the U.S. Department of Energy and the CC was heavily abused during the 1960s as indicated by Center for Applied Biodiversity Science, Conservation dates on discarded soda cans and batteries. The entrance to CC International. I thank the owners for access to caves on their is currently gated with a steel pipe and solid door. The property (PC, FC) and for allowing access to state-owned gate should be re-designed because it alters air flow and pre- caves that required crossing private property. Finally, I thank vents access to bats. The expected aquatic fauna in CC was J.R. Holsinger for encouraging me to undertake this project. absent. The large, deep phreatic pools of CC should have yield- ed populations of troglobitic amphipods and isopods, which REFERENCES were present in the early 1970s (A. Padgett, pers. comm.). A cave fish (Typhlichthes subterraneus) had been collected from Aley, T. & Aley., C., 1991, Delineation and hazard area map- CC in 1971 and was deposited in the University of Georgia ping of areas contributing water to significant caves, in Natural History Museum, Athens. This study detected petrole- Cave Management Symposium, Bowling Green, , um in the cave stream and mud sediments. A truck carrying p. 1-10. petroleum products wrecked on Highway Rt. 136 in the late Barr, T.C., Jr., 1963a, Ecological classification of cavernicoles: 1970s on Lookout Mountain above CC (A. Padgett, pers. Cave Notes, v. 5, p. 9-12. comm.). It is probable that this spill accounts for the lack of Barr, T.C., Jr., 1963b, Studies on the cavernicole Ptomaphagus aquatic cave fauna in CC and illustrates the fragile nature of of the United States (Coleoptera: Catopidae): Psyche, v. 70, cave ecosystems and the difficulty in protecting them. p. 50-58. The remaining caves studied did not appear to have any Barr, T.C., Jr., 1964, Non-troglobitic Carabidae (Coleoptera) significant management concerns. Periodic monitoring should from caves in the United States: Coleopterist’s Bulletin, v. occur to assess long term trends in cave faunal populations. 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96 • Journal of Cave and Karst Studies, December 2001 BUHLMANN

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Muma, M.H., 1946, North American Agelenidae of the genus Reeves, W.K., Jensen, J.B., & Ozier, J.C., 2000, New faunal Coras Simon; American Museum Novitates, no. 1329, p. and fungal records from caves in Georgia, USA: Journal of 20. Cave and Karst Studies, v. 62, p. 169-179. Nicholas, B.C., 1960, Checklist of the macroscopic troglobitic Roth, V.D., 1988, Linyphiidae of America North of Mexico. organisms of the United States: American Midland Checklists, synonymy and literature cited: Gainesville, Naturalist, v. 64, p. 123-160. Florida, American Arachnological Society. Park, O., 1947, Observations on Batrisodes (Coleoptera: Roth, V.D., 1993, Spider genera of North America with keys to Pselaphidae), with particular reference of the American families and genera, and a guide to literature: Gainesville, species east of the Rocky Mountains: Chicago Academy of Florida, American Arachnological Society. Sciences Bulletin, v. 8, p. 45-132. Schultz, G.A., 1970, Descriptions of new subspecies of Peck, S.B., 1973, A systematic revision and the evolutionary Ligidium elrodi (Packard) comb. nov. and notes on other biology of the Ptomaphagus (Adelops) beetles of North isopod crustaceans from caves in North America America (Coleoptera; Leiodidae; Catopinae), with empha- (Oniscoidea): American Midland Naturalist, v. 84, p. 36- sis on cave-inhabiting species: Bulletin of the Museum of 45. Comparative Zoology, v. 145, p. 29-162. Shear, W.A., 1972, Studies in the milliped order Chordeumida Peck, S.B., 1998, A summary of diversity and distribution of (Diplopoda): A revision of the family Cleidogonidae and a the obligate cave-inhabiting faunas of the United States and reclassification of the order Chordeumida in the New Canada: Journal of Cave and Karst Studies, v. 60, p. 18-26. World: Bulletin of the Museum of Comparative Zoology, v. Peck, S.B. & Lewis, J.J., 1978, Zoogeography and evolution of 144, p. 151-352. the subterranean invertebrate faunas of Illinois and south- Wynn, A.H., Highton, R., & Jacobs, J.F., 1988, A new species eastern Missouri: Bulletin of the National Speleological of rock-crevice dwelling Plethodon from Pigeon Mountain, Society, v. 40, p. 39-63. Georgia: Herpetological Review, v. 44, p. 135-143. Platnick, N.I., 1999, A revision of the Appalachian spider Genus Liocranoides (Araneae: Tengellidae): American Museum Novitates, no. 3285, p. 9-13.

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